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Body Size and Flight Distance in Stingless Bees (Hymenoptera: Meliponini): Inference of Flight Range and Possible Ecological Implications

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Body Size and Flight Distance in Stingless Bees (Hymenoptera: Meliponini): Inference of Flight Range and Possible Ecological Implications BODY SIZE AND FLIGHT RANGE IN STINGLESS BEES 563 BODY SIZE AND FLIGHT DISTANCE IN STINGLESS BEES (HYMENOPTERA: MELIPONINI): INFERENCE OF FLIGHT RANGE AND POSSIBLE ECOLOGICAL IMPLICATIONS ARAÚJO, E. D.,1 COSTA, M.,2 CHAUD-NETTO, J.2 and FOWLER, H. G.3 1Grupo de Pesquisa em Comportamento Animal, ITP, UNIT, CEP 49032-490, Aracaju, SE, Brazil 2Departamento de Biologia, Instituto de Biociências, UNESP, Av. 24-A, 1515, C.P. 199, CEP 130506-900, Rio Claro, SP, Brazil 3Departamento de Ecologia, Instituto de Biociências, UNESP. Av. 24-A, 1515, C.P. 199, CEP 130506-900, Rio Claro, SP, Brazil Correspondence to: Edilson D. Araújo, Laboratório de Zoologia, ITP, UNIT, CEP 49032-490, Aracaju, SE, Brazil, e-mail: [email protected] Received February 4, 2003 – Accepted April 30, 2003 – Distributed August 31, 2004 (With 2 figures) ABSTRACT We examined the spatial implications of maximum flight distance for several species of stingless bees. Data suggested that maximum flight distance in Meliponini is a function of body size, especially gen- eralized wing size, which can be estimated through principal component analysis. For six species of stingless bees, flight distances and generalized wing sizes were highly correlated (r = 0.938). This in- dicates that species of Meliponini occupy an effectively larger area as body size increases, which has important implications in the spatial dynamics of local populations restricted to forest fragments. We also used the fitted linear regression model to estimate the maximum flight distance for 12 other species of Meliponini. The results of this research may provide insights for future studies of biological conservation. Key words: flight distance, local populations, morphometry, multivariate analysis. RESUMO Tamanho do corpo em Meliponini (Hymenoptera: Apidae): inferência do raio de vôo e possíveis implicações ecológicas Neste trabalho analisamos as implicações espaciais da distância máxima de vôo para algumas espécies de meliponíneos. Os dados sugerem que a distância máxima de vôo em meliponíneos está relacionada ao tamanho do corpo, especialmente ao tamanho generalizado das asas, que pode ser estimado utilizando análises de componentes principais. Uma análise utilizando seis espécies de meliponíneos evidenciou que o tamanho generalizado das asas está fortemente correlacionado à distância de vôo (r = 0,938). Isso sugere que espécies de meliponíneos ocupam área efetivamente maior quanto maior for o tamanho do corpo, trazendo importantes implicações para a dinâmica espacial de populações locais restritas a áreas fragmentadas. Neste trabalho, também utilizamos um modelo de regressão linear a fim de estimar as distâncias máximas de vôo para 12 outras espécies de meliponíneos. Esta pesquisa fornece subsídios para futuros estudos relacionados à conservação da biodiversidade. Palavras-chave: distância de vôo, populações locais, morfometria, análise multivariada. Braz. J. Biol., 64(3B): 563-568, 2004 564 ARAÚJO, E. D. et al. INTRODUCTION an orthogonal analysis to transform it into a new non-correlated variable set, whose variances decline Meliponini are eusocial bees which are in each principal component, in such a way that the abundant in subtropical and tropical regions of the first principal component explains the major part world, and comprise one of the most diverse and of the variance present in the original characters important insect groups. The stingless bees have (Reis et al., 1988). A logarithmic data transformation an important ecological role, especially in the is generally used to correct for nonlinearity produced pollination of a large fraction (40% to 90%) of the by the allometry among differing morphological plants forming Brazilian forests (Kerr et al., 1994). characters (Gould, 1966; Neff & Marcus, 1980). Data of maxima flight distances of stingless Species studied and their identification codes were: bee workers are important for tropical ecosystems (Pb) Plebeia droryana Friese (1900), (Ts) Trigona (Roubik & Aluja, 1983) and may even be used to spinipes Fabricius (1793), (Mc) Melipona com- formulate models of population dynamics of pressipes Fabricius (1804), (Mq) Melipona different species. Studies of flight distances, in quadrifasciata Lepeletier (1836), (Mm) Melipona conjunction with other studies of local populations, marginata Lepeletier (1836), and (Cc) can generate important inferences on migration, Cephalotrigona capitata Smith (1854), which were colonization, forest fragmentation, and biodiversity chosen because flight distances for these bees can conservation. easily be found in the literature. Flight distances have been studied for only a Morphometric characters were measured using few stingless bee species and generally utilize indirect the criteria of Cunha (1973, 1991) and are given methods of release and recapture (Roubik, 1989). in Table 1, as well as the respective score for each Apparently, there is no directional interference in of the six species studied. flight activities, and the number of marked honey Flight distance data were taken from publi- bees (Apis mellifera) collected declines linearly in cations which used mark-recapture methods. Roubik relation to distance from the colonies (Paranhos et & Aluja (1983) used a magnetic recapture method al., 1997). There is an apparent positive correlation and through regression analysis concluded that the between body size, especially wing area, and flight maximum flight distance for workers of distances (Casey et al., 1985; Byrne et al., 1988). Cephalotrigona capitata was 1,650 m. In Melipona Schwarz (1948) suggested that, because they marginata, maximum flight distance is 800 m. strengthen wing stability, wing hamuli in greater (Wille, 1983). Kerr (1987) recorded a flight dis- number are associated with larger flight capacity. tance of 2,000 m for Melipona quadrifasciata; 840 Worker bee size reflects an adaptation to m for Trigona spinipes; 2,470 m for Melipona environmental conditions (Ruttner, 1988). A major compressipes; and 540 m for Plebeia droryana. part of the morphological variation in Meliponini Two principal component analyses were per- occurs independently of phylogeny due to the fact that, formed. One analysis used generalized body size, for social bees, worker body size has been generally employing all the morphometric variables; the other considered as an adaptation to foraging activity and used only wing variables. First principal component floral resource exploitation (Roubik & Ackerman, scores, for each of these analyses, together with the 1987; Baumgartner & Roubik, 1989). About 75.5% flight distances for each species, were used to form of body size variation in Meliponini corresponds to two new matrices which were examined with a linear adaptive factors associated with resource exploitation regression analysis (Zar, 1999). A third regression (Pignata & Diniz-Filho, 1996). Here we examine the analysis was performed using log values of the spatial implications of maximum flight distance for number of wing hamuli for each of the six species several species of stingless bees. studied in relation to their respective flight distances. Upon fitting of the linear regression model, MATERIAL AND METHODS a new matrix was created with data from the 11 characters of the wing (characters number 18 to 28 Variation in worker body size of six species of Table 1) of 12 other species of Meliponini. This of Meliponini was studied using principal com- new covariance matrix, following the same pattern ponent analysis (PCA) based on covariance matrices. as in the previous analysis, was converted into 11 The PCA uses the original variable set and performs log-transformed characters and subjected to a PCA. Braz. J. Biol., 64(3B): 563-568, 2004 BODY SIZE AND FLIGHT RANGE IN STINGLESS BEES 565 TABLE 1 Measured morphological characters and their scores in the six stingless bee species analyzed (P. droryana – Pd, T. spinipes – Ts, M. compressipes – Mc, M. quadrifasciata – Mq, M. marginata – Mm, and C. capitata – Cc). N Character Pd Ts Mc Mq Mm Cc 1 Length of the antennal flagellus 1.125 1.825 3.100 2.835 0.975 2.425 2 Scape length 0.525 0.925 1.500 1.400 1.025 1.225 3 Mandibular length 0.525 0.925 1.575 1.550 1.025 1.175 4 Inter-malar distance 0.800 1.250 1.625 1.500 1.125 1.400 5 Glossa length 0.950 1.800 2.765 2.475 1.875 2.125 Distance between the central ocellus and the clypeus 6 0.850 1.550 2.500 2.375 1.625 2.200 along the frontal line 7 Inter-ocular distance at the central ocellus 0.625 1.325 2.275 1.800 1.375 1.450 8 Inter-ocular distance along the frontal-clypeus suture 0.400 0.500 1.175 1.225 0.925 0.950 9 Clipeal length 0.200 0.350 0.575 0.475 0.325 0.450 10 Inter-alveolar distance 0.300 0.450 0.700 0.600 0.500 0.525 11 Distance between lateral ocelli 0.225 0.525 0.725 0.750 0.400 0.625 Distance between the lateral ocellus and the 12 0.950 1.800 2.800 2.125 1.625 0.875 compound eye 13 Femur length 1.250 2.800 3.750 3.220 2.150 1.225 14 Tibial length 0.425 0.875 1.425 1.125 0.850 0.500 15 Maximum width of the tibia 0.650 1.125 1.950 1.625 1.175 0.475 16 Basitarsal length 0.275 0.575 1.000 0.825 0.550 0.275 17 Maximum width of the basitarsus 3.600 6.900 9.200 7.800 5.400 7.900 18 Maximum length of the fore wing 1.350 2.475 3.250 2.900 1.925 2.905 19 Maximum width of the fore wing 0.925 1.500 1.800 1.300 1.000 1.900 20 M nerve length 0.100 0.075 0.300 0.225 0.150 0.100 21 Rs nerve length 1.000 1.750 2.870 2.425 1.725 2.200 22 Anal nerve length 0.950 1.800 2.590 2.475 1.650 2.150 23 M + cubital nerve length 0.750 1.350
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